The extinct family Serphitidae in Late Vendean () Michael S. Engel, Vincent Perrichot

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Number 10J

The extinct wasp family Serphitidae in Late Cretaceous Vendean amber (Hymenoptera) Michael S. Engel and Vincent Perrichot

December 1, 2014 Lawrence, Kansas, USA ISSN 1946-0279 (online) paleo.ku.edu/contributions Paleontological Contributions

December 1, 2014 Number 10J

THE EXTINCT WASP FAMILY SERPHITIDAE IN LATE CRETACEOUS VENDEAN AMBER (HYMENOPTERA)

Michael S. Engel1 and Vincent Perrichot2,3*

1Division of Entomology (Paleoentomology), and Department of Ecology & Evolutionary Biology, University of Kansas, 1501 Crestline Drive – Suite 140, Lawrence, Kansas 66045, USA, [email protected], 2CNRS UMR 6118 Géosciences & Observatoire des Sciences de l’Univers de Rennes, Université Rennes 1, 263 avenue du Général Leclerc, 35042 Rennes, France, and 3University of Kansas Biodiversity Institute, Division of Ento- mology (Paleoentomology), Lawrence, Kansas 66045, USA, [email protected]

ABSTRACT A new species of the extinct genus Serphites Brues (Proctotrupomorpha: Bipetiolarida: Serphitidae) is described from two individuals preserved in Late Cretaceous ( to Santonian) amber from Vendée, northwestern France. Serphites fan- nyae n. sp., is distinguished from its congeners and brief comments are made on the significance of finding a serphitid wasp in Vendean amber as well as potential character polarities in the family Serphitidae. Keywords: Insecta, , Proctotrupomorpha, Bipetiolarida, Cretaceous, France RÉSUMÉ Une nouvelle espèce du genre fossile Serphites Brues (Proctotrupomorpha: Bipetiolarida: Serphitidae) est décrite d’après deux spécimens préservés dans l’ambre crétacé supérieur (Cénomanien à Santonien) de Vendée, dans le nord-ouest de la France. Serphites fannyae n. sp., est comparée à ses congénères et la présence d’un Serphitide dans l’ambre de Vendée est brièvement commentée ainsi que la possible polarité de caractères pour la famille Serphitidae. Mots-clés: Insecta, Apocrita, Proctotrupomorpha, Bipetiolarida, Crétacé, France

INTRODUCTION & Engel, 2011), Spathiopterygidae of the Diaprioidea (Engel & There are several families of Hymenoptera which are diagnostic others, 2013), and Falsiformicidae of the Chrysidoidea (Rasnistyn, for the Cretaceous. Each of these families is known almost exclu- 1975; Perrichot & others, 2014). In all cases but Falsiformicidae, sively from amber and typically ranges from the Early Cretaceous individuals are relatively small and their virtual exclusivity to amber through the latter stages of the Mesozoic, and obviously none have is assuredly a representation of the suitability of this medium for been hitherto discovered in the Paleogene. These families include preserving minute organisms with sufficient fidelity. Not surprisingly the Maimetshidae of the Trigonalyoidea (Perrichot & others, 2011), for clades of Mesozoic parasitoids, hosts of these lineages remain Serphitidae of the Serphitoidea (Brues, 1937; Kozlov & Rasnitsyn, unknown and entirely speculative thereby hindering any conclusions 1979; McKellar & Engel, 2011a; Engel, Grimaldi, & Ortega-Blanco, as to whether the occurrence of their victims greatly influences their 2011; Ortega-Blanco & others, 2011a), Alavarommatidae and Gal- presence and abundance in specific deposits, although some kind lorommatidae of the (Engel & Grimaldi, 2007; Gibson, Read, & Huber, 2007; Ortega-Blanco & others, 2011b), of association must assuredly exist. Syninclusions have provided to Stigmaphronidae and Radiophronidae of the Ceraphronoidea (Ko- date no insight into possible host taxa and all that can be surmised zlov, 1975; Engel & Grimaldi, 2009; Ortega-Blanco, Rasnitsyn, & has come from their phylogenetic association with related lineages Delclòs, 2010; McKellar & Engel, 2011b; Ortega-Blanco, Delclòs, (e.g., Engel & Grimaldi, 2009). *Corresponding author.

© 2014, The University of Kansas, Paleontological Institute. | urn:lsid:zoobank.org:pub:88C67F1B-11B3-4A35-9FEF-ACCE4E3A8C08 Engel and Perrichot— New Serphites from French amber 47

The family Serphitidae was first recognized by Brues (1937) based road between La Garnache and Challans, in the department of Ven- on a couple of male inclusions in Campanian amber from Alberta, dée, northwestern France. In a preliminary account of this amber Canada. As can be deduced from the name, Brues considered the deposit, Perrichot and others (2013) suggested a Santonian age but family to be related to the Serphidae (a synonym of Proctotrupidae), contradictory data have been found since then, so an uncertainty although he did note several possible associations and placed particular remains on the exact age and until new elements are found we can- emphasis on the two-segmented petiole, an otherwise rare feature not discriminate between Middle Cenomanian, Turonian, or Early among the Hymenoptera. Subsequent to Brues’s (1937) account there Santonian (≈ 97–85 Ma), as discussed by Perrichot and Néraudeau remained almost no work on the family for nearly half a century. In (2014: 10A in this volume). Morphological terminology and the 1979, Kozlov and Rasnitsyn described several new serphitids, includ- format for the description generally follows that of Engel, Grimaldi, ing two new genera, from the Cenomanian and Santonian amber of and Ortega-Blanco (2011) and McKellar and Engel (2011a). Photo- and noted a number of traits shared between serphitids and graphs were taken with a Canon 5D Mark II digital camera attached the , particularly the form of the two-segmented to a Leica MZ APO stereomicroscope, and stacks of images taken petiole (later the combined clade was named Bipetiolarida). Indeed, at different depths of field were merged using Helicon Focus 5.3 Kozlov and Rasnitsyn (1979) suggested merging the two as subfami- (HeliconSoft Ltd) to obtain sharpness throughout the entire images. lies within a single family. Again, the group remained dormant until Metrics were taken with an ocular micrometer set on an Olympus recently several additional taxa were added based on a revision of the SZX-12 stereomicroscope. original Canadian fauna (McKellar & Engel, 2011a) as well as mate- rial from the Albian of Spain (Ortega-Blanco & others, 2011a) and SYSTEMATIC PALEONTOLOGY Turonian of New Jersey (Engel, Grimaldi, & Ortega-Blanco, 2011). Family SERPHITIDAE Brues, 1937 A review of the fauna from is presently underway Genus SERPHITES Brues, 1937 by one of us. Recently, two serphitids have been recovered in the Type species.—Serphites paradoxus Brues, 1937, p. 33, fig. 5A. Cenomanian–Santonian amber of Vendée in northwestern France. Included species.— Presently 12 species included in the genus Herein is provided a brief account of this material. (see the key below). MATERIAL AND METHODS Comment.—The emended generic diagnosis by McKellar and Engel (2011a) is followed here and requires no modification to The material comprises two specimens. The first individual (Figs. incorporate the present species. J1.1, J2.2) is preserved in a small piece of clear yellow amber with one true fly (Diptera: Dolichopodidae: Microphorinae: Microphorites Key to species of Serphites magaliae Perrichot & Engel – see issue 10G in this volume) as a 1. Pterostigma isosceles ...... 2 syninclusion. The specimen is mostly complete, although the gaster Pterostigma equilateral ...... 3 is missing beyond the first gastral segment and portions of some legs 2. Body small, 1.3 mm long in male; 1st petiolar segment twice as are missing at the amber surface. Some areas are difficult to observe long as 2nd petiolar segment, the latter flattened dorsally; gaster owing to deformities in the amber surface (particularly near the with lateral carina between tergites and sternites [Campanian, metasoma), a large blackened bit of debris to the specimen’s right, Canada] ...... S. paradoxus Brues and the position of M. magaliae masking the specimen’s face. The Body large, 3.0 mm long in female; 1st petiolar segment 3× as second serphitid (Figs. J1.2, J2.1) is similarly preserved but rests long as 2nd petiolar segment, the latter not flattened dorsally; alongside a darkened fracture plane rendering it impossible to see the gaster longer than mesosoma, without lateral carina [Santonian, specimen in dorsal-oblique view from the left. This orientation also Taimyr] ...... S. gigas Kozlov & Rasnitsyn resulted in the wings being less than ideally preserved (hind wings 3. 1st petiolar segment longitudinally striate ...... 4 present, right forewing largely destroyed except for extreme base; left 1st petiolar segment not longitudinally striate [Cenomanian– forewing present but mixed into fracture plane making interpreta- Santonian, France] ...... S. fannyae n. sp. tion challenging) and portions of the dorsum of the prothorax and 4. 1st petiolar segment not rimmed anteriorly ...... 5 head are obscured, including a direct facial view. Nonetheless, this 1st petiolar segment rimmed anteriorly [Early Albian, Spain] ..... is the most complete individual and because the head is twisted, ...... S. lamiak Ortega-Blanco, Delclòs, Peñalver, & Engel gives the best view of the opened mouthparts along with antennal 5. Gaster shorter than mesosoma ...... 6 structure. This is also the only specimen with a complete metasoma Gaster as long as or slightly longer than mesosoma ...... 9 (the sting is exposed along with the short associated styli), and it 6. Metatarsus shorter than metafemur; metabasitarsus shorter than was duly selected as the holotype. combined length of remaining tarsomeres ...... 7 The amber pieces were slightly polished and subsequently em- Metatarsus as long as or longer than metafemur; metabasitarsus bedded in blocks of epoxy resin which were again polished on all as long as combined length of remaining tarsomeres ...... 8 sides. This facilitated optimal viewing of the specimens, although 7. Body small, female 1.10 mm long, male 1.45 mm long; forewing the ideal vantage for particular structures was still not possible given vein Rs not tubular, straight; 1st petiolar segment 2.7× as long the placement of syninclusions or other impurities with the amber. as 2nd petiolar segment; 1st and 2nd gastral segments shorter The amber was collected in 2002 by Magali Weigandt and Fanny than combined length of remaining segments [Turonian, New Dupé from a deposit exposed briefly during work along the D32 Jersey] ...... S. raritanensis Engel & Grimaldi 48 Paleontological Contributions, number 10J

10. Body length 1.5 mm; forewing vein Rs nebulous; 1st petiolar segment twice as long as 2nd petiolar segment; 1st and 2nd gastral segments shorter than combined length of remaining segments [Campanian, Canada] ...... S. hynemani McKellar & Engel Body length 2.0 mm; forewing vein Rs tubular; 1st petiolar segment 2.4× as long as 2nd petiolar segment; 1st and 2nd gastral segments shorter than combined length of remaining segments [Santonian, Taimyr] ...... S. dux Kozlov & Rasnitsyn Body length 2.1 mm; forewing vein Rs tubular; 1st petiolar segment 3× as long as 2nd petiolar segment; 1st and 2nd gastral segments longer than combined length of remaining segments [Campanian, Canada] ...... S. bruesi McKellar & Engel SERPHITES FANNYAE new species Figures J1–J2 Type material.—Holotype female IGR.GAR-26 (Figs. J1.2, J2.1), ex coll. Dupé, and paratype male IGR.GAR-106b (Figs. J1.1, J2.2), ex coll. Weigandt; in Late Cretaceous (Middle Cenomanian to Early Santonian, ≈ 97–85 Ma) Vendean amber. Both specimens are deposited in the Geological Department and Museum of the University Rennes 1, France. Type locality.—La Robinière, departmental road D32, about 2.5 km south-west of La Garnache, Vendée, France. Etymology.—The specific epithet is a matronym honoring Fanny Dupé, who collected one of the amber pieces containing the type series. Diagnosis.—Total body length under 2 mm (differs from S. navesinkae at ca. 2.6 mm and S. silban and S. gigas at ca. 3 mm); trochantelli short (long in S. lamiak, extremely short and superficially Figure J1. Photographs of Serphites fannyae n. sp., in Late Cretaceous (Cenoma- absent in S. gigas and S. dux); metatarsus very slightly longer than nian–Santonian) amber of Vendée, NW France; 1, paratype male IGR.GAR- metafemur (about as long as in S. silban and S. gigas); metabasitarsus 106b; 2, holotype female IGR.GAR-26. Scale bars = 0.25 mm. distinctly shorter than combined length of remaining tarsomeres, about two-thirds length of remainder of metatarsus (as long as or Body larger, female 2.6 mm long; forewing vein Rs tubular, nearly so in S. lamiak and S. raritanensis, about twice in S. dux); curved apically; 1st petiolar segment 3× as long as 2nd first petiolar segment not rimmed anteriorly (rimmed inS. lamiak); petiolar segment; 1st and 2nd gastral segments longer than second petiolar segment not flattened dorsally (flattened dorsally in combined length of remaining segments [Campanian, S. paradoxus); first petiolar segment not longitudinally striate (stri- Canada] ...... McKellar & Engel S. kuzminae ate in most species); first petiolar segment about twice the length 8. Body length 2.6 mm, female with nine antennomeres; left of second petiolar segment (more than twice, 2.7–3× as long in S. mandible with lower tooth slightly longer than upper tooth; raritanensis and S. kuzminae, respectively); and gaster longer than metatarsus longer than metafemur; 1st petiolar segment 2.5× mesosoma (shorter in S. raritanensis, as long as in S. dux). as long as 2nd petiolar segment; 1st and 2nd gastral segments Description.—Female. Total body length 1.63 mm; forewing length shorter than combined length of remaining segments [Turonian, 0.82 mm. Integument apparently dark brown (although many areas New Jersey] ...... S. navesinkae Engel & Grimaldi with artificial metallic silver sheen owing to microscopic separation Body length 2.95 mm; left mandible with lower and upper from amber causing reflections), with scattered, minute, fine setae teeth equal in size; metatarsus as long as metafemur; 1st petiolar throughout body, such setae particularly numerous on tibiae and segment twice as long as 2nd petiolar segment; 1st and 2nd tarsi. Head not greatly enlarged but with relatively sizeable gena, gena gastral segments longer than combined length of remaining only slightly narrower than compound eye; antenna 10-segmented; segments [Early Albian, Spain] ...... scape barely extending half-way to vertex, with length approximately S. silban Ortega-Blanco, Delclòs, Peñalver, & Engel three times the greatest width (0.14 mm long); flagellum with relative 9. Body very small, 0.9 mm long; basal flagellomere about 1/3 as lengths of articles 1.0, 0.8, 0.9, 1.2, 1.4, 1.4, 1.5, 2.0; first flagellar long as apical one; right mandible with middle tooth longest article chalice-like in outline, second article nearly square in outline, of three teeth, left mandible with two equal teeth [Campanian, following articles almost rectangular in outline, broader than long, Canada ...... S. pygmaeus McKellar & Engel fifth article broadest flagellomere; apical flagellar article terminating Body length at least 1.5 mm; right and left mandibles with lower in rounded acute point; right mandible tridentate, lower two teeth tooth longest ...... 8 longest and most prominent, uppermost tooth distinctly shorter, Engel and Perrichot— New Serphites from French amber 49

Figure J2. Line drawings of Serphites fannyae n. sp., Late Cretaceous (Cenomanian–Santonian) amber of Vendée, NW France. 1, habitus of holotype female IGR.GAR-26; 2, lateral left habitus of paratype male IGR.GAR-106b. apparently one-half length of midtooth, lowermost tooth longest, long, terga and sterna apparently smooth with minute punctures at only slightly longer than midtooth, all teeth with sharply pointed setal bases, discs of terga and sterna with several scattered, suberect apices; left mandible bidentate, both teeth long with sharp apices, setae, sterna not concave, instead slightly convex on discs, segments lower tooth distinctly longer than upper tooth. Mesosoma 0.53 mm generally of same size except apically larger; sting exposed, relatively in length, mostly obscured by preservation and with dorsal surface short, scarcely longer than associated sheaths and only somewhat not visible. Forewing with numerous microtrichia, slightly longer longer than an individual gastral segment (although the sting may setae forming continuous fringe along wing margins; venation typi- not be fully exserted), sting simple, apparently without preapical cal for species of the genus (see McKellar & Engel, 2011a; Engel, dorsal notch. Grimaldi, & Ortega-Blanco, 2011); C not fused to Sc+R, forming a Male. Very similar but specimen incomplete, with most of distinct costal cell, C not pigmented (perhaps due to preservation); gaster missing so total length in life is unknown. Body length as pterostigma massive, triangular, approximately equilateral, uniformly preserved 1.28 mm. Forewing length 0.84 mm. Mesosomal length pigmented and sclerotized; r-rs arising from pterostigmal midlength, 0.52 mm. Antenna 9-segmented, which suggests it is a male rather about as long as wide; Rs strongly pigmented, not tubular, straight, than a female according to the sexual dimorphism in the number reaching wing margin; Rs+M absent; M+Cu, basal vein, and 1Cu of antennomeres of Serphites. tubular, other abscissae of Cu and M nebulose, apicalmost abscissa of M reaching wing margin, that of Cu disappearing shortly before DISCUSSION margin; 1A tubular proximally, becoming nebulous well before 1Cu. The discovery of a serphitid wasp in Vendean amber, as well as Hind wing with numerous microtrichia, with short setae forming a species of the microphorine genus Microphorites Hennig, 1971 marginal fringe, with only C+Sc+R along anterior margin and with (Perrichot & Engel, 2014: 10G in this volume), is in accordance three hamuli apically. Legs thin, although not especially long, with with the Cretaceous age of this amber even though the precise scattered, minute, fine setae; trochantelli short; femora not especially layer is unknown within the Cenomanian–Santonian interval. Both swollen; meso- and metatibiae with two short, thin apical spurs Microphorites and, as noted above, the Serphitidae are classical Cre- (often difficult to discern from surrounding setae), single protibial taceous amber taxa, and both became extinct sometime by the end spur; tarsi pentamerous; metabasitarsus two-thirds length of remain- of the Mesozoic. The new species can be easily assigned to Serphites ing tarsomeres combined; pretarsal claws simple and short, arolium based on the eight-segmented female flagellum (six-segmented in large. Metasoma bipetiolate; first petiolar segment not rimmed Jubaserphites McKellar & Engel, 2011), absence of distinctively anteriorly (as in S. raritanensis), about twice the length of second elongate and numerous setae on the vertex (present in Jubaserphites), petiolar segment, petiolar segments apparently smooth and without pronotum large and extending back to the tegula (not reaching distinct sculpturing; gaster slightly longer than mesosoma, 0.63 mm tegula in Microserphites Kozlov & Rasnitsyn, 1979), pterostigma 50 Paleontological Contributions, number 10J large, well defined, and sclerotized (indistinct inMicroserphites ), rvie NOVAMBRE 2 (both to D. Néraudeau, Univ. Rennes 1). This first petiolar segment twice length of second petiolar segment (less work is a contribution of the Division of Entomology, University of than twice length in Aposerphites Kozlov & Rasnitsyn, 1979), and Kansas Biodiversity Institute. lateral ocellus nearly touching compound eye (distinctly separated from compound eye margin in Aposerphites). Serphites is the most diverse and widespread of the four genera in Serphitidae and may REFERENCES ultimately prove to be paraphyletic with respect to one of the other Brues, C. T. 1937. Superfamilies Ichneumonoidea, Serphoidea, and Chal- genera, thereby requiring its segregation into multiple groups. For cidoidea. University of Toronto Studies, Geological Series 40:27–44. the time being serphitids remain too poorly known to speculate Engel, M. S., & D. A. Grimaldi. 2007. 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Peñalver, & M. S. Engel. 2011a. Ser- ACKNOWLEDGEMENTS phitid wasps in Early Cretaceous amber from Spain (Hymenoptera: We are grateful to Fanny Dupé and Magali Weigandt who col- Serphitidae). Cretaceous Research 32(2):143–154, DOI: 10.1016/j. lected the amber pieces containing the specimens described above cretres.2010.11.004. and who made possible their study; to Didier Néraudeau who Ortega-Blanco, J., E. Peñalver, X. Delclòs, & M. S. Engel. 2011b. False fairy wasps in Early Cretaceous amber from Spain (Hymenoptera: My- assisted with the initial screening of the amber; and to Alexandr marommatoidea). Palaeontology 54(3):511–523, DOI: 10.1111/j.1475- Rasnitsyn and a anonymous reviewer for their valuable comments 4983.2011.01049.x. on the first draft of the manuscript. Partial support was provided Perrichot, V., & D. Néraudeau. 2014. Introduction to thematic volume by French National Research Agency grant nº BLAN07-1-184190 "Fossil in Late Cretaceous Vendean amber (northwestern (project AMBRACE) and CNRS-INSU grant through project Inter- France)". Paleontological Contributions 10A:1–4. Engel and Perrichot— New Serphites from French amber 51

Perrichot, V., & M. S. Engel. 2014. Youngest occurrence of the genus Mi- France. 6th International Congress on Fossil Insects, Arthropods, and crophorites (Diptera: Dolichopodidae): A new species in Late Cretaceous Amber. Byblos, Lebanon, 14th–18th April 2013, abstract volume p. 49. Vendean amber. Paleontological Contributions 10G:30–33. Perrichot, V., M. S. Engel, A. Nel, J. Ortega-Blanco, X. Delclòs, C. Soriano, Perrichot, V., J. Ortega-Blanco, R. C. McKellar, X. Delclòs, D. Azar, A. V. Lohrmann, & P. Tafforeau. 2014. "False ants" from the Cretaceous (Hymenoptera: Chrysidoidea: Falsiformicidae). 11th meeting of Hyme- Nel, P. Tafforeau, & M. S. Engel. 2011. New and revised maimetshid nopterologists in Stuttgart. Mitteilungen des Entomologischen Vereins wasps from Cretaceous (Hymenoptera, Maimetshidae). ZooKeys Stuttgart 49:26. 130:421–453, DOI: 10.3897/zookeys.130.1453. Rasnitsyn, A. P. 1975. Hymenoptera Apocrita of Mesozoic. Trudy Pale- Perrichot, V., D. Néraudeau, V. Girard, A. Nel, Y. Nohra, S. Saint Martin, ontologicheskogo Instituta, Akademii Nauk SSSR [Transactions of the J.-P. Saint Martin, A. R. Schmidt, & F. Dupé. 2013. Santonian Vendeen Paleontological Institute, Academy of Sciences of the USSR] 147:1–134 amber: large amounts of data from a small sample in north-western [In Russian].